Dtsch. Entomol. Z. 61 (2) 2014, 105–119 | DOI 10.3897/dez.61.8142

museum für naturkunde

A taxonomic revision of Parallelodemas Faust from South China (Coleoptera, , ), with notes on sexually dimorphic characters

Jens Prena1, Runzhi Zhang1

1 Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Beijing 100101, P. R. China http://zoobank.org/B449C54C-A8A8-4B00-826A-ED41A3BE9D53

Corresponding author: Runzhi Zhang ([email protected])

Abstract Received 19 June 2014 Accepted 19 August 2014 Nine of Parallelodemas Faust are reported from China. In addition to the pre- Published 17 September 2014 viously recorded P. impar Voss, we found P. docile Faust, P. vicinum Faust and the fol- lowing six species newly described herein: P. dimetans sp. n., P. feae sp. n., P. petilum Academic editor: sp. n., P. plumosum sp. n., P. setifrons sp. n. and P. tumens sp. n. Parallelodemas tarsale James Liebherr Voss from Java is transferred to Lepidomyctides Yoshihara and Morimoto (comb. n.). Lectotypes are designated for P. docile, P. imperfectum Faust, P. perfectum Faust, P. tar- dum Faust and P. vicinum. Several morphological modifications with unknown functions Key Words are documented, illustrated and discussed. Males of some species have special setae on rostrum, antennal scape and mesotarsus. A medially notched epistome apparently oc- curs in both sexes but seems to wear off in females, probably during the preparation of sexual dimorphism oviposition sites. The mandible is unusual in having a convex, edentate inner face and exodontous mandible incisor-like structures on the outer face. Oriental Region

Introduction Material and methods

Among the currently accepted 548 genera of baridine wee- Our study is based primarily on specimens of the Institute vils, Parallelodemas Faust is notable for having numerous of Zoology of the Chinese Academy of Sciences (IZCAS), morphological peculiarities. Faust (1894) noticed modified which were collected during the past 60 years. Their col- setae on the male mesotarsus of two of his five species and lecting data are transcribed herein to Pinyin (original Chi- a loss of the two distal tarsites in another [actually present nese spelling is given for primary types) and we provide but minute]. Voss (1941) commented on the basally bifur- the unique IZCAS database identifiers in square bracket cate prosternum of female P. impar Voss, and Marshall behind the collecting date. Additional specimens were (1945) and Morimoto and Yoshihara (1996) on apparently studied from the following collections: Forschungsmuse- abducent mandibles. To this one may add the frequently um Alexander Koenig, Bonn, Germany (AKMB); Natural worn female epistome and marked sexual dimorphism of History Museum, London, UK (BMNH); Bernice P. Bish- further body parts, such as rostrum, antennal scape and eye, op Museum, Honolulu, Hawaii (BPBM); Canadian Muse- in at least some species. However, this interesting genus is um of Nature, Ottawa, Canada (CMNC); Naturhistorisches taxonomically challenging because of great morphological Museum Basel, Switzerland (NHMB); Jens Prena person- similarity of the species, pronounced sexual dimorphism al collection, Rostock, Germany (JPPC); Museo civico and scarcity of material in collections. China is presently di storia naturale Giacomo Doria, Genoa, Italy (MSNG); the only country with noteworthy collections of Parallelo- Senckenberg Naturforschendes Museum, Frankfurt a. demas. In this paper, we revise the Chinese species and Main, Germany (SFFM); Senckenberg Naturhistorische document taxonomically important characters of the genus. Sammlungen, Dresden, Germany (SNSD); Zhejiang A & F

Copyright Prena J. & Zhang R. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which per- mits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 106 Jens Prena & Runzhi Zhang: A taxonomic revision of Parallelodemas Faust from South China...

University, Zhejiang, China (ZAFU) and Zoological Insti- tute of the Russian Academy of Sciences, St. Petersburg, Russia (ZIN). The above codens are used to refer to col- lections in the text. A total of 154 specimens was exam- ined. Faust returned approximately half of the type series of his newly described species to L. Fea (the owner) and shared the rest with L. v. Heyden. Whenever possible, we designated as lectotype a male from the material returned to Fea and ignored the unpublished designations in Faust’s retained material, which were made by museum staff sub- sequent to the acquisition of his collection. If not provided on the label, coordinates of collecting sites were determined with GoogleEarth. Approximate collecting sites of Fea’s specimens, from his 1885–89 journey to Myanmar, were estimated based on his travel itinerary (Vinciguerra 1890; Fea 1896). Descriptions are structured hierarchically, i.e., constant generic characters are not repeated in the descriptions of species. We found no evidence for a species with a sexually dimorphic epi- stome. However, the female epistome apparently wears off so we described it only for the male. Measurements of length were taken with an ocular micrometer in a ste- reomicroscope. Total length was measured from the ante- rior margin of eye to the abdominal apex in dorsal view. Length-width ratios of the penis are approximate values which do not take into account curvature. Illustrations were prepared from images taken with a Micropublisher 5.0 RTV digital CCD camera mounted on a Zeiss Ste- REO Discovery V12 or a Canon EOS 650D on a Leica DM2500 compound microscope. Aldus Freehand was used for vector graphics and Adobe Photoshop for pix- Figure 1. Parallelodemas docile, dorsal habitus (length 5.2 mm). el-based artwork. All genitalia illustrations were prepared from specimens collected in China. Explanations for Parallelodemas from certain grass- and sedge-associated, morphological terms can be found at http://weevil.info/ primarily Palaeotropical Baridinae with similarly slender glossary--characters. Our weevil classification fol- rostrum and elongate body. The latter complex includes lows Oberprieler et al. (2014) except that we maintain Eumycterus Schönherr, 1838; Trephognathus Marshall, Baridinae and as separate subfamilies. 1945; Neosharpia Hoffmann, 1956; Caenobaris Nasreddi- nov, 1980; Lepidomyctides Yoshihara & Morimoto, 1994 and several species currently placed in other genera. Misplaced species. Yoshihara and Morimoto (1994) recognized that Parallelodemas tarsale Voss, 1937 is a Genus Parallelodemas Faust species near Eumycterus and Lepidomyctides but they had very little material of those genera. We studied five Parallelodemas Faust, 1894: 306. Type species Parallelo- Oriental species near P. tarsale (BPBM, IZCAS, SNSD, demas perfectum Faust, by subsequent designation ZIN) and several species of Caenobaris, Eumycterus, (Morimoto and Yoshihara 1996). Gender originally Neosharpia and Trephognathus Marshall from Africa, female, changed to neuter because of the grammat- Central Asia and India. While Eumycterus and its prob- ical gender of the Ancient Greek base word δέμας able synonym Trephognathus can be distinguished by (Alonso-Zarazaga and Lyal 1999). vertically moving mandibles (Marshall 1945; Korotyaev 2002), we were unable to recognize or to confirm the ge- Diagnosis. Species of Parallelodemas can be recognized neric limits of the remaining species. We transfer here P. by characteristically elongate body (Fig. 1), medially tarsale to Lepidomyctides in the widest sense, as Lepido- notched epistome (which often is worn in females) and myctides tarsalis (Voss), new combination. exodontous mandibles with evenly convex inner face Redescription. Habitus: Total length 3.0–7.8 mm, width (Fig. 3). Superficially, they resemble species of the conod- 0.8–2.2 mm; body slender subcylindrical (Fig. 1); integu- erine subtribe Phaenomerina (see Morimoto 1961), but ment black or brown, appendages and ventrites sometimes those have incrassate, ventrally dentate femora and larger rufous; vestiture uniform or locally condensed to short eyes. The characters on the mandible and epistome separate vittae, setae either simple, squamiform, scalloped, deeply

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Figure 3. Rostrum of P. plumosum, male (left) and female (right), dorsal view, showing apex with epistome and mandibles. Figure 2. Rostrum of P. impar, male (left) and female (right), lateral view. agreeing with modal arrangement of subfamily (Zherikhin and Gratshev 1995). Abdomen: Ventrites unmodified, not split or plumose. Head: Subspherical, contour often slight- or indistinctly sexually dimorphic. Sclerolepidia small to ly warped at rostral base; eyes large, slightly encroaching medium-sized, densely packed, digitate. Stridulatory devic- on rostrum, bulging or flush with head contour, dorsally es absent. Male genitalia and associated structures: Tergite separated by width of rostrum at base; frontal fovea small VII without plectra for stridulation; tergite VIII shorter than to moderate; rostrum moderately elongate and slender, wide, distally without transverse carina; sternite VIII later- slightly curved, female with apical portion slightly inflated ally with sclerotized pyriform area, medially desclerotized; (Fig. 2); epistome produced and more or less notched, often sternite IX variously strongly curved, distal prongs narrow- worn off in females (Fig. 3); scrobe laterally descending, ly to widely diverging but always symmetrical; penis dor- inserted between midlength and apical fourth; fu- soventrally depressed, longer than basal apodemes; inter- nicle with 7 desmomeres; club compact, spindle-shaped, nal sac extending approximately to midlength of apodemes basal article approximately as long as remainder of club, when inverted, with sclerite at insertion of duct or with pig- not annexed to distal desmomere; mandibles with apparent- mented flagellum; tegmen with ring dorsally closed, basal ly abducent movement (away from center line and slightly apodeme obsolete, parameroidal lobes developed. Female ventrad), outer face with 1 large and 1–2 small secondary genitalia and associated structures: Tergite VII longer than teeth, inner face convex and without teeth (Fig. 3). Protho- wide, without transverse carina, setal vestiture squamiform rax: Barrel-shaped, elongate, nearly as wide as elytra. Ante- basally and piliform distally, plectra for stridulation ab- rior margin of pronotum not projected over frons, subapical sent; sternite VIII distally forked into weakly sclerotized, constriction absent; basal margin bisinuous to accommo- widely dilated, U-shaped arms; hemisternite pigmented, date projecting base of elytron; postoccular lobe feeble or stylus 1.9–2.2× as long as wide, distal setae half as long absent. Prosternum without median channel, rarely slightly as stylus; bursa pouch-like, half as long as vagina; sperma- depressed in front of coxae; subapical constriction slight to theca sclerotized, collum short, often somewhat bulbous, moderate; basal lobe partially projected over mesosternum, ramus inserted on outer face of collum (facing away from with basal margin bifurcate or (rarely) truncate. Pterotho- cornu), rudimentary to long; spermathecal duct unpigment- rax: Mesoscutellum visible, trapezoid to subquadrate. Me- ed, at most slightly longer than spermatheca, inserted dis- sosternum unmodified. Mesepimeron smaller and narrower tally in bursa. Legs: Procoxae separated by less than 1/3 than mesepisternum, ascending between pronotum and el- diameter of coxa; pro- and mesofemora clavate, hindfemur ytron and visible in dorsal view. Metasternum medially de- less expanded and often partially sulcate ventrally; tibiae pressed in male, flat or convex in female. Elytra: Elongate, straight to curved (depending on ventral profile of femur), sides subparallel, apices rounded individually, humerus de- ventrodistal spine spiniform, robust and large on pro- and veloped, subapical callus feeble or absent, base at interstriae mesotibiae but somewhat smaller on metatibia; tarsus with 3–6 slightly depressed and somewhat projected; striae 10, 5 segments, third with anterior margin faintly to deeply ex- narrow but distinct, strial punctures not or slightly affect- cised, fifth long to greatly reduced, claws flat and basally ing edge of interstriae, strial setae absent; interstriae flat, fused, or miniaturized and medially fused to single blade. punctate to transversely rugose, interstrial setae uniform Diversity and distribution. With the six new species or heterogeneous, modified setae restricted to basal and described in this study, Parallelodemas includes now a submedian vittae if present. Hindwings: Fully developed, total of twelve. The scarce material gives an unrepre- length-width ratio 3.4–3.7, fore margin basally concave, sentative picture of the distributional ranges of individ- anal lobe moderate, hind margin with setal fringe; venation ual species. Species of Parallelodemas have been found

dez.pensoft.net 108 Jens Prena & Runzhi Zhang: A taxonomic revision of Parallelodemas Faust from South China... in China, India, Laos, Malaysia, Myanmar, Taiwan and we could obtain neither direct field observations nor fresh Vietnam. Their distribution is primarily Oriental but sev- specimens for scanning the abductor and adductor tendons. eral reach the Palaeartic part of China, northward up to Eye. While almost all Baridinae have eyes that are Shaanxi and Zhejiang. flush with the head contour, they are protruded in sever- Biology. The host of Parallelodemas apparently al Parallelodemas species. The eyes of male P. setifrons is unknown. One specimen of P. docile is labeled as being protrude more than those of females (Fig. 8), but the dor- taken from Buttontree, Anogeissus acuminata (Roxburgh sal and ventral distance between them and their circum- ex Candolle) Guillemin et al. (Combretaceae). Other spec- ference are not affected. The increased eye surface affords imens were swept from low vegetation. Females with ful- more facets in the male but facet diameter is the same. ly developed eggs occur from late April to early June. Leg. Several Parallelodemas species have large, devi- Sexual dimorphism. Rostrum. Species of Paral- ant setae on the mesotarsus which crowd toward the distal lelodemas exhibit marked sexual dimorphism of char- (outer) half. These setae are arranged asymmetrically on acters on the rostrum. Females generally have a longer the fifth (claw-bearing) tarsite and are much larger and and smoother rostrum than males, with a more basal- more numerous in males than in females (Figs 6, 7). The ly inserted antenna and slightly inflated apical portion individual tarsites, in particular the fifth, are often more (Fig. 2). Gender-related differences in the basal width elongate in males than in females. Males often have faint- of the rostrum (Fig. 8) are apparent but often inconspic- ly thicker pro- and mesofemora than females. uous. The ventral side of the rostrum is setose in male P. Tergites. Like in other Baridinae, the eighth tergite is impar, P. petilum and P. plumosum. However, the most developed in males but internalized in females. Because striking difference, the length and shape of the epistome, the distal external tergite protrudes beyond the elytral may be acquired secondarily rather than being truly sex- apex well enough to expose the suture between the sev- ually dimorphic. Nearly all examined males have a pro- enth and eighth tergites in males, this character is very jected, medially notched epistome, whereas it is almost useful for sexing specimens. always short and truncate in females (Fig. 3). However, Ventrites. The male metaventrite is medially de- the presence of projected epistomes in some female P. pressed and, together with the first abdominal ventrite, feae, P. impar, P. imperfectum and P. setifrons suggests may have less setae than the female’s. Voss (1941) men- a secondary loss, probably through abrasion during the tioned a sexually dimorphic basal process on the proster- preparation of oviposition sites, because the distally di- num of P. impar, but he either had a mixed series or his vergent mandibles afford no protection of the epistome observation was incorrect. as in other weevils. However, this needs to be confirmed with freshly emerged specimens and field observations. Antenna. Males generally have a longer scape than fe- males (usually as long as the funicle). The distal margins of the male scape can be setose, such as in P. impar (Fig. 2). Mandible. At a first glance, it appears as if Parallelode- mas has swapped the left with the right mandible or rotated them by 180 degrees (Fig. 3). The inner face not only lacks any trace of incisors, it also is evenly convex from base to apex and seems therefore dysfunctional. Moreover, the out- er face is concave and armed with two apparently ordinary incisors, just like the inner mandibular face of most baridine weevils. However, three landmarks on the mandible leave no doubt that the seemingly abducent mandibular movement Figures 4–7. Left mesotarsus of Parallelodemas, dorsal view: evolved by reversing the function of the abductor and ad- 4. P. docile; 5. P. imperfectum; 6. P. vicinum, male; 7. P. vici- ductor tendons rather than by rotating the mandible, a trend num, female. Not to scale. seen in some weevils with a very slender rostrum (Mar- shall 1945): (1) The dorsal and ventral mandibular sockets (preartis and postartis) are formed and located as in other Baridinae; (2) the mandibular setae are located on the out- er face (between the basal incisors); and (3) the pharyngeal process is attached to the inner basal angle of the mandible. From this it follows that the incisors on the outer face are secondarily evolved structures and analogous to the inner incisors of other weevils. Morimoto and Yoshihara (1996) suggested an inversion of the mandibular movement from adducent to abducent. The laterally deeply excised mandib- Figure 8. Parallelodemas setifrons, dorsal view of head show- ular articulation and widely divergent mandibles in many ing sexually dimorphic protrusion of eyes and rostral width (left mounted specimens support this conclusion. Unfortunately, male, right female).

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Key to the species of Parallelodemas found in China

1 Thoracic sternites with plumose or deeply split setae...... 2 – Thoracic sternites with simple or scalloped, indistinctly split setae...... 5 2 Thoracic sternites with bi- and trifid setae, plumose setae absent...... 3 – Thoracic sternites with plumose setae...... 4 3 Eyes slightly bulging; male rostrum glabrous ventrally; total length 3.6–4.3 mm...... P. dimetans – Eyes distinctly bulging; male rostrum hirsute ventrally; total length 3.0–3.7 mm...... P. petilum 4 Frons at most with a few setae; eyes slightly bulging in both sexes; penis with apex gradually rounded (Fig. 14), internal sac with harpoon-shaped sclerite; total length 3.6–5.4 mm...... P. plumosum – Frons hirsute; eyes distinctly bulging in male; penis with apex distinctly pointed and produced (Fig. 15), internal sac medially with small, fennel-seed-like sclerite and near duct with sclerotized area with spinose lateral projection; total length 3.0–3.9 mm...... P. setifrons 5 Tarsite 3 large, anterior margin slightly to moderately excised (Figs 4, 5); claws miniaturized, not longer than fifth tar- somere wide...... 6 – Tarsite 3 small, anterior margin deeply excised beyond middle (Figs 6, 7); claws longer...... 7 6 Tarsite 5 inserted near middle and exceeding anterior margin of tarsite 3 by half its own length (Fig. 4); male rostrum distinctly shorter than pronotum; total length 4.5–6.2 mm...... P. docile – Tarsite 5 minute, inserted in distal fifth and barely exceeding margin of tarsite 3 (Fig. 5); male rostrum as long as pro- notum; total length 5.4–6.4 mm; Myanmar...... P. imperfectum 7 Eyes bulging; profemur slender, subdistally gradually converging; male rostrum ventrally setose; male antenna inserted in apical third of rostrum...... P. impar – Eyes nearly flush with head contour; profemur clavate, subdistally noticeably constricted in lateral view; male rostrum ventrally glabrous; male antenna inserted in mid-third of rostrum...... 8 8 Male mesotarsus without clavate setae; male antenna with scape glabrous along proximal edge; prosternum in front of coxa often tumescent in lateral view; aedeagus apically lancet-shaped...... 9 – Male mesotarsus with clavate setae; male antenna with scape setose along proximal edge; prosternum in front of coxa usually gradually sloping in lateral view; aedeagus subparallel...... 10 9 Rostrum shorter (male <0.95×, female 1.04× length of pronotum); male antenna inserted more distally (prorostrum 0.33× length of rostrum); total length 5.4–6.6 mm; Myanmar...... P. tardum – Rostrum longer (male >1.05×, female 1.20× length of pronotum); male antenna inserted more basally (prorostrum 0.42× length of rostrum); total length 3.8–4.8 mm; China (Guizhou)...... P. tumens 10 Profemur hirsute ventrally; metepisternum with squamiform setae rather evenly distributed; total length 4.9–6.5 mm...... P. feae – Profemur squamose ventrally; metepisternum with squamiform setae increasingly larger and denser in distal section; total length 6.4–7.8 mm...... 11 11 Metepisternum distally with imbricate squamiform setae; male profemur moderately expanded ventrally; penis with apex trunctate (Fig. 20)...... P. vicinum – Metepisternum distally with more widely spaced squamiform setae; male profemur strongly clavate, ventrally almost angular; penis with apex narrowly rounded (as P. feae, Fig. 19); Myanmar...... P. perfectum

Parallelodemas dimetans Prena & Zhang, sp. n. frons and base of rostrum with some recumbent setae; http://zoobank.org/596CADD7-085B-42F5-A76D-7DF7F6B4CDC2 male rostrum 1.07× as long as pronotum, ventrally with- out setae, prorostrum 0.41–0.42× rostral length, slightly Diagnosis. This small species can be recognized by the spatulate and apically diverging in dorsal view, epistome presence of slender, bi- or trifid setae on the thoracic ven- slightly notched, antennal scape with a few long setae, trites and ventrally glabrous male rostrum. Parallelode- club 1.7× as long as wide; female rostrum 1.02–1.11× as mas petilum is the only other known species with such long as pronotum, prorostrum 0.47–0.51× rostral length; setae but the male rostrum is hirsute. prosternum gradually sloping in front of coxa, basal lobe Description. Length 3.6–4.3 mm, width 1.0–1.2 mm; notched; all femora hirsute ventrally; tarsus with tarsite 3 integument dark brown, antenna, tarsus, apex of female relatively small and excised to basal third, tarsite 5 short- rostrum and often other parts of leg brown or rufous; er than 2+3 and distinctly protruding beyond anterior ventral side and pygidium with simple, slender setae, margin of 3, male mesotarsus with long, distally pointed thoracic sterna also with bi- and some trifid setae, basic setae; penis 2.5× as long as wide, apex roundly narrowed vestiture of fine setae on pronotum and elytron,- some and slightly projected medially (Fig. 12), internal sac with what larger white setae at base of elytral interstria 3 and harpoon-like sclerite and small sclerotized area near duct; postmedially on interstriae 3 and 4; eyes slightly bulging; spermatheca with nodulus and ramus short (Fig. 21).

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Distribution. The species is known only from China with moderately long, clavate, outward directed setae; (Guizhou). penis 7× as long as wide, apex triangularly narrowed (as Material examined. Holotype: China, Guizhou P. plumosum, Fig. 14), with sclerotized flagellum almost Province, Suiyang County, Kuankuoshui [宽阔水] Nat- as long as penis and apodemes combined; spermatheca ural Reserve, Xiangguang Village [香广山村], 27.9798 with collum and nodulus forming poorly differentiated N 107.1661 E, 1550 m, 4.vi.2010 [#1500871], Wang bulbous unit, ramus as long as wide (Fig. 22). Zhiliang [王志良], male (IZCAS). Paratypes (3 males, Distribution. The species (in the strict sense) is known 10 females): CHINA. Guizhou: Suiyang, Kuankuoshui from the Chinese province Shaanxi. Natural Reserve, Forestry Station Holiday Center, 1206 Material examined. Holotype: China, Shaanxi Prov- m, 5.vi.2010 [#1500879], Wang Zhiliang (IZCAS 1); ince, Huoditang forest farm [火地塘林场], 1559 m, Suiyang, Kuankuoshui Natural Reserve, Gongtong ravine, 33.4343 N 108.4480 E, 14.viii.2013, Jiang Chunyan [姜 6.vi.2010 [#1500880], Nie Cuie (IZCAS 1); Suiyang, 春燕], male, dissected, #1941160 (IZCAS). Paratypes (1 Kuankuoshui Natural Reserve, Jinzi Village, 3.vi.2010 male, 1 female): same data as holotype, male, #1941162 [#1500876–78], Wang Zhiliang (IZCAS 3); Suiyang, (IZCAS 1), female, #1941161 (IZCAS 1). Other material: Kuankuoshui Natural Reserve, Xiangguang Village, CHINA. Sichuan: Mount E’mei, Jiulao cave, 1800–1900 4.vi.2010 [#1500872–74], 8.vi.2010 [#1500875], Wang m, 1.viii.1957 [#1500865], Lu Youcai, male (IZCAS 1). Zhiliang (IZCAS 4); Suiyang, Kuankuoshui Natural Re- VIETNAM. Cao Bang Prov.: Mount Pia Oac, 7.vi.2011, serve, 3.vi.2010 [#1500867–68], 4.vi.2010 [#1500869], S. Lingafelter, 2 males, 1 female (IZCAS 1, JPPC 2). Liu Wangang (IZCAS 3); Suiyang, Kuankuoshui Natural Etymology. The name is a Latin adjective for slender Reserve, Baishao ravine, 8.vi.2010 [#1500882], Nie Cuie or gaunt. (IZCAS 1). Etymology. The name is a participle presence active of dimeto (=to delimit, to mark-off; Latin). Parallelodemas plumosum Prena & Zhang, sp. n. http://zoobank.org/1D130D15-F6BC-4F76-AC26-78B10E438383

Parallelodemas petilum Prena & Zhang, sp. n. Diagnosis. This species can be distinguished from P. seti- http://zoobank.org/ED12DF0F-C531-4943-B5CA-14C8AECAAB0A frons, the other known species with plumose setae, by glabrous frons and slightly protruding eyes in both sexes. Diagnosis. Our material includes seven small specimens Parallelodemas dimetans and P. petilum have at most tri- from three sites (3 Shaanxi, 1 Sichuan, 3 Vietnam), which fid setae. form a close-nit complex of probably three species. They Description. Length 3.6–5.4 mm, width 1.0–1.4 mm; have bulging eyes, thoracic ventrites with bifid setae, a integument black, antenna and tarsi dark brown; ventral ventrally setose male rostrum and a slender penis with side with plumose setae, pygidium with bifid setae, pro- very long flagellum. Differences occur in the apical shape notum and elytron with basic vestiture of very fine setae, of the penis (Shaanxi – triangular; Sichuan – narrowly elytron also with short vittae of plumose white setae at rounded; Vietnam – slightly projected) and the first ab- base of interstria 3 and postmedially on interstriae 3–5; dominal ventrite (Vietnamese specimens with a pair of eyes very slightly bulging; frons and base of rostrum with tubercles between the metacoxae). We describe the three a few recumbent setae; male rostrum 0.94–1.06× as long specimens from Shaanxi as P. petilum and informally as- as pronotum, ventrally without setae, prorostrum 0.40– sign to this complex the four others. Similar species are 0.43× rostral length and slightly spatulate in dorsal view, P. dimetans (with nearly flush eyes) andP. setifrons (with epistome moderately notched, antennal scape with a few plumose setae). long setae, club 1.8× as long as wide; female rostrum Description. Length 3.0–3.7 mm, width 0.8–1.0 mm; 1.00–1.04× as long as pronotum, prorostrum 0.49–0.51× integument dark brown to nearly black, appendages par- rostral length; prosternum gradually sloping in front of tially light brown; meso- and metathoracic sternites with coxa, basal lobe truncate; pro- and mesofemora hirsute bi- and trifid setae, pronotum and elytron with basic ves- ventrally; tarsus with tarsite 3 moderately large and ex- titure of very fine setae; eyes bulging; frons and base of cised to basal third, tarsite 5 nearly as long as 1 and dis- rostrum with recumbent setae; male rostrum 0.93–1.12× tinctly protruding beyond anterior margin of 3, with long, as long as pronotum, ventrally with long setae, pro- distally unmodified setae in both sexes; penis 3× as long rostrum 0.39–0.40× rostral length, slightly spatulate and as wide, roundly narrowed to subtriangular tip (Fig. 14), apically diverging in dorsal view, epistome moderately internal sac with harpoon-like sclerite; spermatheca with deeply notched, antennal scape with a few long setae, nodulus short and thick, ramus as long as wide (Fig. 24). club 2.6× as long as wide; female rostrum 1.14× as long Distribution. The species occurs in China (Fujian, as pronotum, prorostrum 0.50× rostral length; prosternum Hainan) and Taiwan. gradually sloping in front of coxa, basal lobe moderately Material examined. Holotype: China, Fujian Prov- notched; all femora hirsute ventrally; tarsus with tarsite 3 ince, Jianyang [建阳], Chong’an Xin Village [崇安星 excised to basal third, tarsite 5 as long as 2 and distinctly 村], Sangang [三港], 740 m, 27.7489 N 117.6831 E, Pu protruding beyond anterior margin of 3, male mesotarsus Fuji [蒲富基], 14.v.1965, male, dissected, #1799537 (IZ-

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CAS). Paratypes (13 males, 6 females): CHINA. Fujian: with nodulus as long as wide and perpendicular to long Jianyang, Chong’an Xin Village, Qili Bridge, 12.vii.1963 axis of collum, ramus as long as nodulus. [#1500864], Zhang Youwei (IZCAS 1); Jianyang, Distribution. The species is known from China (Fujian,­ Chong’an Xin Village, Sangang, 29.v.1960 [#1799132], Guangdong, Guizhou, Hunan). 4.vi.1960 [#1500862], 7.vi.1960 [#1799540], Jiang Material examined. Holotype: China, Fujian, Jian­ Shengqiao (IZCAS 3), 6.vii.1960 [#1799529], Pu yang, Chong’an Xin Village [崇安星村], Sangang [三 Fuji (IZCAS 1), Zhang Yiran (IZCAS 1), 14.v.1965 港], 27.7489 N 117.6831 E, 740 m, 29.v.1960, Zuo Yong [#1500863]; Jianyang, Chong’an Tongmuguan Guan- [左永], male, dissected, #1799533 (IZCAS). Paratypes ping, 22.v.1960 [#1799136], Jiang Shengqiao (IZCAS (5 males, 14 females): CHINA. Fujian: Dazhulan, 1); Mount Wuyi, Guadun, 3.vii.1982 [#1500861], Jiang Shaowu, 19.–25.v.1945 [#1941180] (IZCAS 1); Kuatun Fan (IZCAS 1). Hainan: Qiongzhong, Mount Wuzhi, [=Guatun], 5.v.1938 (2×), 5.vi.1938, J. Klapperich ancient plant path, 29.vii.2009 [#1500852], Liang Hong- (AKMB 3); Jianyang, Huangkeng Changba, 8.iv.1960 bin (IACAS 1), 30.xi.2009 [#1879661–62, #1879689], [#1500856], Jiang Shengqiao (IZCAS 1); Jianyang, Zhang Kuiyan & Lin Meiying (IZCAS 3); Qiongzhong, Huangkeng Guilin, 5.iv.1960 [#1799536, #1799549], Mount Wuzhi, main peak, 28.xi.2009 [#1879660], Huang 14.iv.1960 [#1799139, #1799158], Jiang Shengqiao Lina (IZCAS 1); Qiongzhong, Mount Wuzhi, Shuiman (IZCAS 4); Jianyang, 17.v.1965 [#1500855] (IZCAS 1); Village, 30.vii.2009 [#1500853–54], Huang Xinglei (IZ- Jianyang, Sangang, Chong’an Xin Village, 14.v.1960 CAS 2); Mount Diaoluo, Holiday Village, 12.xii.2008 [#1799135], Pu Fuji (IZCAS 1), 29.v.1960 [#1799533], [#1500851], Wang Zhiliang (IZCAS 1). TAIWAN. Zuo Yong (IZCAS 1). Guangdong: Ruyuan, Huangling, Nanjing: Wushe, 3.v.1983 (2×), 10.v.1983, H. Townes Mount Xiaohuang, 18.vii.2008 [#1799020], Li Yingchao (CMNC 3). (IZCAS 1); Nanling, Ruyang Natural Reserve Station, Etymology. The name is a Latin adjective meaning 19.vii.2008 [#1799016], Li Yingchao (IZCAS 1). plumed, or with feathers. Guizhou: Kuankuoshui Natural Reserve, Gongtong ravine, 7.vi.2010 [#1500866], Nie Cuie (IZCAS 1); Suiyang, Kuankuoshui Natural Reserve, Jinzi Village, Parallelodemas setifrons Prena & Zhang, sp. n. 3.vi.2010 [#1500859], Wang Zhiliang (IZCAS 1); http://zoobank.org/3F48D0AF-32EE-47DE-A838-8B37AECD3F7B Suiyang, Kuankuoshui Natural Reserve, Xiangguang Village, Suiyang, 4.vi.2010 [#1500857–58], Wang Diagnosis. Useful characters for identification are Zhiliang (IZCAS 2); Suiyang, Kuankuoshui Natural plumose setae, hirsute frons and sexually dimorphic eyes. Reserve, 4.vi.2010 [#1500860], Liu Fanggang (IZCAS Parallelodemas petilum is similar but has at most trifid 1). Hunan: Yizhang, Mount Mang, forest park west gate, setae and a ventrally hirsute male rostrum. 17.vii.2008 [#1799015], Li Yingchao (IZCAS). Description. Length 3.0–3.9 mm, width 0.9–1.0 mm; Etymology. The name is a Latin noun in apposition integument dark brown, antenna, legs and apex of rostrum composed of seta and frons. light brown; meso- and metathoracic sterna with plumose setae (condensed on metepisternum), prosternum, ab- dominal ventrites and pygidium mostly with bi- and trifid Parallelodemas docile Faust setae, pronotum and elytron with basic vestiture of fine setae, elytron also with short vittae of white, moderately Parallelodemas docilis [sic] Faust 1894: 309. Lectoype wide, simple or bifid setae at base and postmedially on male, designated here, labeled “Carin Cheba/ 900-1100 interstriae 3–5; eyes distinctly (male) or slightly (female) m/ L. Fea V XII-88”, “docilis/ Faust”, “P./ docilis/ sp. bulging; frons and base of rostrum with recumbent setae; n.”, “Parallelodemas/ docilis/ ♂ sp. n.”, “SYNTYPUS/ male rostrum 0.84–0.90× as long as pronotum, ventral- Parallelodemas/ docilis/ Faust, 1894”, “Museo Civico/ ly with long setae, prorostrum 0.43–0.48× rostral length di Genova”, “Lectotype/ Parallelodemas docile/ Faust and slightly spatulate in dorsal view, epistome slightly (Prena & Zhang/ design. 2014)” (MSNG). Paralecto- notched, antennal scape without long setae, club 1.4–2.4× types 6: Carin Cheba, Myanmar (MSNG 3, SFFM 1, as long as wide (depending on length of rostrum); female SNSD 2). rostrum 0.84–1.00× as long as pronotum, prorostrum 0.49–0.57× rostral length; prosternum gradually sloping Diagnosis. The large, subcordate third and relatively in front of coxa, basal lobe notched; all femora hirsute short fifth tarsite (Fig. 4) are diagnostic for this species. ventrally; tarsus with tarsite 3 relatively small and excised Parallelodemas imperfectum Faust, the only other spe- to basal third, tarsite 5 nearly as long as 1 and distinctly cies with enlarged third tarsite, has an extremely min- protruding beyond anterior margin of 3, male mesotarsus iaturized fifth tarsite (incorrectly stated to be absent by with moderately long, clavate, outward directed setae; Faust 1894). Other useful characters are the short male penis 2.5× as long as wide, with small, apically rounded rostrum and presence of dense vestiture on the distal three projection (Fig. 15), internal sac medially with small, fen- fourths of the metepisternum. nel-seed-like sclerite and with sclerotized area with spi- Redescription. Length 4.5–6.2 mm, width 1.1–1.6 nose lateral projection near duct; spermatheca (Fig. 24) mm; integument black, teneral specimens with ventrites

dez.pensoft.net 112 Jens Prena & Runzhi Zhang: A taxonomic revision of Parallelodemas Faust from South China... and legs partially dark rufous; ventral side with undi- legs and ventrites brown; ventral side with undivided vided setae, basic vestiture inconspicuous on pronotum setae, basic vestiture on pronotum and elytron absent, and elytron, imbricate white squamiform setae at base imbricate yellowish white squamiform setae at base of elytral interstria 3, postmedially on interstriae 3–5, on of elytral interstria 3, postmedially on interstriae 3–4, dorsal apex of mesepimeron, distal 3/4 of metepisternum, ventrally and laterally on thorax and abdomen (includ- flank of prosternum, ventral face of pro- and mesofem- ing basolateral angles of pronotum) and on ventral face ora, dorsal face of metafemur and occasionally on ba- of pro- and mesofemora; eyes flush with head contour; solateral angles of pronotum; eyes flush with head con- frons and base of rostrum glabrous; male rostrum 0.94× tour; frons and base of rostrum glabrous; male rostrum as long as pronotum, ventrally without setae, prorostrum 0.78–0.83× as long as pronotum, ventrally without setae, 0.35× rostral length and slightly spatulate in dorsal view, prorostrum 0.38–0.41× rostral length and subcylindrical epistome moderately notched in 1 female [worn in oth- in dorsal view, epistome very slightly notched, antennal er specimens including male], antennal scape ventrally scape ventrally without long setae, club 1.6× as long as without long setae, club 1.4× as long as wide; female ros- wide; female rostrum 1.02–1.13× as long as pronotum, trum 1.02–1.07× as long as pronotum, prorostrum 0.50× prorostrum 0.50–0.53× rostral length; prosternum gradu- rostral length; prosternum gradually sloping in front of ally sloping in front of coxa, basal lobe slightly notched; coxa, basal lobe slightly notched; pro- and mesofemora pro- and mesofemora ventrally with recumbent squami- ventrally with recumbent squamiform setae; tarsus with form setae; tarsus with tarsite 3 moderately large, sub- tarsite 3 moderately enlarged, subcordate, anterior mar- cordate and excised to middle, tarsite 5 as long as 3 and gin straight and only indistinctly excised, tarsite 5 minute only moderately protruding beyond anterior margin of 3 (Fig. 5); penis 3.0× as long as wide, slightly tapering in (Fig. 4), inconspicuously setose in both sexes; penis 2.5× apical third as in P. docile (Fig. 16) but with apex more as long as wide, apex subtriangular (Fig. 16), internal sac pointed, internal sac with Y-shaped sclerite; spermatheca without discernible sclerite; spermatheca with nodulus with nodulus and ramus short (as P. docile, Fig. 25). and ramus short (Fig. 25). Distribution. The species is known from one site in Distribution. The species is known from China (Yun- Myanmar. nan) and Myanmar. Material examined. MYANMAR. Kayin: Karen Hills Material examined. CHINA. Yunnan: Xishuang- [ca. 25–35 km NE of Taungoo, 900–1100 m], V/1888, L. banna Prefecture, Menglun, 12.iv.1994, 13.vi.1994 Fea (MSNG 2, SNSD 1). [#1799526, #1941179], Yang Long (IZCAS 2); Xi- shuangbanna Prefecture, Menglun, Highway G213, 21.xi.2009 [#1500899–901], 22.xi.2009 [#1500902–05], Parallelodemas impar Voss 26.xi.2009 [#1500906–07], Tang Guo & Yao Zhiyuan (IZCAS 9); Xishuangbanna Prefecture, Tropical Plant Parallelodemas impar Voss 1941: 895. Holotype female, Garden, 19.viii.2007 [#1500898], Zheng Guo (IZCAS Tienmushan, China (NKMB). Paratype retained by 1]. MYANMAR. Kaya: Karen Hills, Iadó [=Yahku?], Voss not traceable and probably destroyed (see Weid- XII/1887 (MSNG 2). Kayin: Karen Hills, Leitó [=Leik- ner 1979: 400). tho], V/1888, L. Fea (MSNG 2, SFFM 1, SNSD 1). Diagnosis. The ventrally hirsute male rostrum separates P. impar from all other Chinese species with simple, un- Parallelodemas imperfectum Faust divided setae. Another good character is the absence of wide setae on elytron and metepisternum. Parallelodemas imperfecta [sic] Faust 1894: 310. Lectotype Redescription. Length 4.4–5.8 mm, width 1.0–1.5 male (dissected), designated here, labeled “Carin Che- mm; integument dark brown, antenna, tarsus and apex of ba/ 900-1100 m/ L. Fea V XII-88”, “imperfecta/ Faust”, female rostrum light brown; vestiture consisting of incon- “P./ imperfecta/ sp. n.”, “Parallelodemas/ imperfecta/ ♂ spicuous, evenly distributed, simple, cupreous setae; eyes ♀”, “SYNTYPUS/ Parallelodemas/ imperfecta/ Faust, slightly bulging; frons and base of rostrum with recum- 1894”, “Museo Civico/ di Genova”, “Lectotype/ Paral- bent setae; male rostrum 0.98–1.07× as long as pronotum, lelodemas imperfectum/ Faust (Prena & Zhang/ de- ventrally with long setae, prorostrum 0.28–0.30× rostral sign. 2014)” (MSNG). Paralectotypes 2: Carin Cheba, length and spatulate in dorsal view, epistome deeply Myanmar (MSNG 1, SNSD 1). notched, antennal scape ventrally with long, cupreous setae, club 2.0× as long as wide; female rostrum 1.13– Diagnosis. This species can be recognized by its charac- 1.16× as long as pronotum, prorostrum 0.48–0.49× ros- teristic tarsus: the enlarged third tarsite is barely excised tral length; prosternum gradually sloping in front of coxa, anteriorly and the fifth is greatly reduced (Fig. 5). Par- basal lobe notched; all femora hirsute ventrally; tarsus allelodemas docile has a larger fifth tarsite and is more with tarsite 3 relatively small and excised to basal third, elongate. tarsite 5 as long as 1 and moderately protruding beyond Redescription. Length 5.4–6.4 mm, width 1.5–2.1 anterior margin of 3, male mesotarsus with long, clavate, mm; integument black, antenna and sometimes parts of outward directed setae; penis 3.0× as long as wide, apex

dez.pensoft.net Dtsch. Entomol. Z. 61 (2) 2014, 105–119 113 with narrowly rounded, subtriangular projection (Fig. ventrally with erect squamiform setae; tarsus with tarsite 17), internal sac with thick, tubular, basally curved scler- 3 of moderate size and excised to basal third, tarsite 5 ite; spermatheca with nodulus long and perpendicular to slightly longer than 3 and distinctly protruding beyond long axis of collum, ramus obsolete (Fig. 26). anterior margin of 3, male mesotarsus without specialized Distribution. The species is known from China (Si- setae; penis 2.3× as long as wide, apex lancet-shaped and chuan, Yunnan, Zhejiang) and Laos. The record from broadly rounded (as P. tumens, Fig. 18), internal sac with Guatun in Fujian, by Voss (1956), applies to P. setifrons. 2 small sclerites (hook and paired hook); spermatheca Material examined. CHINA. Sichuan: Wushan Coun- with collum bulbous, ramus short, nodulus obsolete (as ty, Liziping, 2.vii.1993 [#1941174] (IZCAS 1). Yun- P. dimetans, Fig. 21). nan: Xishuangbanna Prefecture, Menghun, 17.v.1958 Distribution. Besides the type series from Myanmar, [#1500892], Meng Xuwu (IZCAS 1). Zhejiang: West we have seen one female from India that might be this Lin’an, Mount Tianmu, 15.6.1957 [#1799534], 28.6.1957 species. [#1500891, #1500893, #1799531] (IZCAS 4), 9.vi.2012 Material examined. INDIA. Sikkim: Gopaldhara, (ZAFU 1); Kaishanlaodian, Mount Tianmu, 23.vi.1998 Rungbong Valley, H. Stevens (BMNH 1). MYANMAR. [#1941175–76] (IZCAS 2); Mount Tianmu (NHMB 1). Kayin: Karen Hills [ca. 25–35 km NE of Taungoo, 900– LAOS. Luang Prabang: Ban Kiukacham env., 19.vi.2009, 1100 m], V/1888, L. Fea (MSNG 3, SNSD 1). M. Geiser & D. Hauck (NHMB 5). Xieng Khouang: Phonsavan to Phu Padaeng, 30/31.v.2009, M. Geiser (NHMB 2). Parallelodemas tumens Prena & Zhang, sp. n. http://zoobank.org/2295F76C-0867-4E0C-980C-1CFB6E45ACE0

Parallelodemas tardum Faust Diagnosis. Besides P. tardum, this is the only known spe- cies without split setae on the metepisternum and without Parallelodemas tarda [sic] Faust 1894: 309. Lectoype male, a modified male mesotarsus. Parallelodemas tumens is designated here, labeled “Carin Cheba/ 900-1100 m/ L. smaller (<5 mm) than P. tardum and has a longer ros- Fea V XII-88”, “tarda/ Faust”, “P./ tarda/ sp. n.”, “Par- trum with a more basally inserted antenna. Differences in allelodemas/ tarda/ ♂ sp. n.”, “SYNTYPUS/ Parallelo- the genitalia are not apparent. Female P. tumens may be demas/ tarda/ Faust, 1894”, “Museo Civico/ di Geno- distinguished from female P. perfectum and P. tardum by va”, “Lectotype/ Parallelodemas tardum/ Faust (Prena smaller body size. All three species have specimens with & Zhang/ design. 2014)” (MSNG). Paralectotypes 3 (2 a more or less tumescent prosternum. males, 1 female), same data (MSNG 2, SNSD 1). Description. Length 3.8–4.8 mm, width 1.1–1.3 mm; integument black, antenna, tarsus, apex of female ros- Diagnosis. Two of the eight known species with undivid- trum and often other parts of leg brown or rufous; ventral ed setae have an unmodified male mesotarsus, i.e., they side and pygidium with undivided setae, basic vestiture lack special setae and the fifth tarsite is not enlarged. One of fine setae on pronotum and elytron, moderately wide is P. tardum described from Myanmar, the other is P. tu- white setae at base of elytral interstria 3 and postmedially mens from China. Parallelodemas tardum is larger than on interstriae 4 and 5, on thoracic flank and basolateral P. tumens (5.4–6.6 mm vs. 3.8–4.8 mm) and has a shorter angles of pronotum; eyes flush with head contour; frons rostrum with a more distally inserted antenna. The fe- and base of rostrum glabrous; male rostrum 1.06–1.08× males may be distinguished by body length. A difference as long as pronotum, ventrally without setae, prorostrum between female P. perfectum and P. tardum is not appar- 0.41–0.42× rostral length, slightly spatulate and apical- ent (each with one known specimen). Female P. feae are ly diverging in dorsal view, epistome short and truncate, very similar but have ventrally hirsute femora. antennal scape glabrous, club 1.8× as long as wide; fe- Redescription. Length 5.4–6.6 mm, width 1.6–1.9 male rostrum 1.20× as long as pronotum, prorostrum mm; integument black; ventral side and pygidium with 0.53× rostral length; prosternum tumescent in front of undivided setae, basic vestiture of fine setae on prono- coxae, basal lobe notched; pro- and mesofemora hirsute tum and elytron, moderately wide white setae at base of ventrally; tarsus with tarsite 3 of moderate size and ex- elytral interstria 3 and postmedially on interstriae 3–5, cised to basal third, tarsite 5 slightly longer than 3 and on metepisternum and basolateral angles of pronotum; distinctly protruding beyond anterior margin of 3, male eyes flush with head contour; frons and base of rostrum mesotarsus without specialized setae; penis 2.5× as long glabrous; male rostrum 0.89–0.94× as long as pronotum, as wide, apex lancet-shaped and broadly rounded (Fig. ventrally without setae, prorostrum 0.32–0.33× rostral 18), internal sac with 2 small sclerites (hook and paired length, apically slightly diverging in dorsal view, epi- hook); spermatheca with nodulus and ramus short (as P. stome short and slightly notched, antennal scape gla- dimetans, Fig. 21). brous, club 1.8× as long as wide; female rostrum 1.04× as Distribution. The species is known from the Chinese long as pronotum, prorostrum 0.50× rostral length; pros- province Guizhou. ternum slightly tumescent in front of coxae (apparently Material examined. Holotype: China, Guizhou Prov- not in female), basal lobe notched; pro- and mesofemora ince, Libo County, Banzhai Village [板寨村], 24.v.1998,

dez.pensoft.net 114 Jens Prena & Runzhi Zhang: A taxonomic revision of Parallelodemas Faust from South China...

Zhang Runzhi [张润志], #1500884, male, dissected (IZ- Mountain Village, 20.v.1999 [#1799153], Han Hong­ CAS). Paratypes (2 males, 1 female): CHINA. Guizhou: xiang (IZCAS 1). Hainan: Jianfeng, Tianchi, 18.iii.1980 Libo County, Banzhai Village, 24.v.1998, Zhang Runzhi [#1500888], Wang Shuyong (IZCAS 1); Ledong County, [#1500883] (IZCAS 1); Yunyi Prefecture, Suiyang Coun- Jianfengling Natural Reserve, 10.iv.1980 [#1500890], ty, Kuankuoshui Natural Reserve, 3.vi.2010 [#1500881], Wang Shuyong (IZCAS 1), 4.v.2007 [#1500889, 4.vi.2010 [#1500870], Nie Cuie (IZCAS 2). #1854471], Ge Deyan (IZCAS 2); Gaotuo Shan [高陀山, Etymology. The name is a participle presence active not located], 17.v.1963 [#1799532], Zhou Yao (IZCAS of tumeo (=to inflate, to distend; Latin). 1). Hunan: Yanling County, Shidu Shennong Valley wa- terfall, 7.vii.2008 [#1799013], Jiao Tianyang (IZCAS 1). Yunnan: Xishuangbanna Prefecture, Meng’a, 11.v.1958 Parallelodemas feae Prena & Zhang, sp. n. [#1799522], Hong Chunpei (IZCAS 1). LAOS. Houa http://zoobank.org/09E6371C-7BC3-4E5F-B117-C66FB9976068 Phan: Phou Pane Mountains, 1350–1500 m, 1.–16. vi.2009, M. Brancucci (NHMB 1). Xieng Khouang: Diagnosis. From other species with undivided setae, P. Phou Sane Mountains, 30 km NE Phonsavan, 1400–1500 feae can be separated by having ventrally hirsute femora m, 10.–30.v.2009, Z. Kraus (NHMB 1). VIETNAM. and nearly flush eyes. Female P. tardum and P. tumens Tinh Vinh Phuc: Tam Dao, 25.v.1995, A. V. Gorochov are very similar but have shorter and wider setae on the (ZIN 1). pro- and mesofemora. Etymology. The name is a patronym honoring the Ital- Description. Length 4.9–6.5 mm, width 1.3–1.8 mm; ian zoologist and artist Leonardo Fea. integument dark brown to black, antenna and tarsus brown; ventral side and pygidium with undivided setae, basic vestiture of fine setae on pronotum and elytron, Parallelodemas vicinum Faust moderately wide white setae at base of elytral interstria 3, postmedially on interstriae 3 and 4 and on thoracic flank; Parallelodemas vicina [sic] Faust 1894: 308. Lectoype eyes very slightly bulging; frons and base of rostrum gla- male, designated here, labeled “Carin Cheba/ 900-1100 brous; male rostrum 1.02–1.20× as long as pronotum, m/ L. Fea V XII-88”, “vicina/ Faust”, “P./ vicina/ sp. ventrally without setae, prorostrum 0.39–0.42× rostral n.”, “Parallelodemas/ vicina/ ♂ sp. n.”, “SYNTYPUS/ length, slightly spatulate and apically diverging in dorsal Parallelodemas/ vicina/ Faust, 1894”, “Museo Civi- view, epistome slightly notched, antennal scape with long co/ di Genova”, “Lectotype/ Parallelodemas vicinum/ setae, club 1.8× as long as wide; female rostrum 1.04– Faust (Prena & Zhang/ design. 2014)” (MSNG). Para- 1.20× as long as pronotum, prorostrum 0.50–0.54× ros- lectotypes 9: Carin Cheba, Carin Ghecu and Teinzó, tral length; prosternum gradually sloping in front of coxa, Myanmar (MSNG 4, SFFM 2, SNSD 3). Lyal and King basal lobe notched; all femora hirsute ventrally; tarsus (1996), elytro-tergal stridulation. with tarsite 3 relatively small and excised to basal third, tarsite 5 nearly as long as 2+3 and distinctly protruding Diagnosis. A generally useful character for recognizing beyond anterior margin of 3, male mesotarsus with clav- P. vicinum is the presence of imbricate squamiform setae ate, outward directed setae; penis 3.5× as long as wide, on the distal half of the metepisternum. Parallelodemas apex roundly narrowed and medially projected (Fig. 19), docile has similar vestiture on the distal two thirds and an internal sac with thick, rod-like sclerite; spermatheca enlarged third tarsite. Small P. vicinum with more widely with nodulus long and perpendicular to collum (Fig. 27). spaced setae on the metepisternum differ from the oth- Distribution. The species is known from China (Fu- erwise very similar P. perfectum by the apically truncate jian, Guangxi, Hainan, Hunan, Yunnan), India, Laos and aedeagus and less curved female rostrum. These two and Vietnam. P. feae, a species with ventrally hirsute femora, are the Material examined. Holotype: China, Hainan Prov- only known species with undivided setae, flush eyes and ince, Jianfeng [尖峰], Tianchi [天池], 18.iii.1980, Wang clavate setae on the male mesotarsus. Shuyong [王书永], #1799133, male, dissected (IZCAS). Redescription. Length 6.4–7.8 mm, width 1.6–2.2 Paratypes (9 males, 10 females): CHINA. Fujian: Jian- mm; integument black, teneral specimens with ventrites gle County, Mount Longxi, 21.i.1991 [#1500887], Yang and legs partially dark rufous; ventral side with undivided Longlong (IZCAS 1), 19.v.1991 [#1500886], Zhang setae, basic vestiture inconspicuous on pronotum and el- Runzhi (IZCAS 1); Jianyang, Chong’an Xin Village ytron, imbricate white squamiform setae at base of elytral Longdu, 7.v.1960 [#1799553], Ma Youcai (IZCAS 1); interstria 3, postmedially on interstriae 3 and 4, on dor- Jianyang, Chong’an Xin Village Sangang, 17.vi.1960 sal apex of mesepimeron, distal half of metepisternum, [#1799530], Jiang Shengqiao (IZCAS 1), 26.vi.1960 flank of prosternum, ventral face of pro- and mesofemora, [#1799542], Zuo Yong (IZCAS 1); Jianyang, Huangkeng dorsal face of metafemur and occasionally on basolateral Dazhulan, 11.vi.1960 [#1500885], Zuo Yong (IZCAS angles of pronotum; eyes flush with head contour; frons 1), 24.vii.1960 [#1799535], Jiang Shengqiao (IZCAS and base of rostrum glabrous; male rostrum 1.06–1.15× 1); Jianyang, Huangkengguilin, 14.iv.1960 [#1799157], as long as pronotum, ventrally without setae, prorostrum Zhang Yiran (IZCAS 1). Guangxi: Jinxiuhuawang 0.36–0.38× rostral length and spatulate in dorsal view,

dez.pensoft.net Dtsch. Entomol. Z. 61 (2) 2014, 105–119 115

Figures 9–11. Male terminalia of P. vicinum: 9. Penis, dorsal view; 10. Tegmen; 11. Sternites 8 and 9, ventral view.

Figures 12–20. Male genitalia of Parallelodemas, apex of pe- epistome very slightly notched, antennal scape with long, nis, dorsal view: 12. P. dimetans; 13. P. petilum; 14. P. plumo- cupreous setae, club 1.6× as long as wide; female rostrum sum; 15. P. setifrons; 16. P. docile; 17. P. impar; 18. P. tumens; 1.14–1.24× as long as pronotum, prorostrum 0.57–0.58× 19. P. feae; 20. P. vicinum. Not to scale. rostral length; prosternum gradually sloping in front of coxa, basal lobe notched; pro- and mesofemora ventrally with slender (male) or squamiform (female) setae; tarsus Carin Ghecú [between Taó and Chialá, II–IV/1888] with tarsite 3 relatively small and excised to basal third, (MSNG 1). Kayin: Karen Hills [ca. 25–35 km NE of tarsite 5 as long as 2+3 and distinctly protruding beyond Taungoo], V/1888, L. Fea (MSNG 3, SFFM 2, SNSD 3). anterior margin of 3, male mesotarsus with moderately long, clavate, outward directed setae; penis 2.5× as long as wide, apex bottle-shaped (Fig. 9), internal sac with Parallelodemas perfectum Faust thick, tubular, basally curved sclerite; spermatheca with nodulus long and usually perpendicular to collum, ramus Parallelodemas perfecta [sic] Faust 1894: 307. Lectoype similarly long (as Fig. 27). male, designated here, labeled “♂ Carin/ Cheba”, “per- Distribution. The species is known from China (Yun- fecta/ Faust”, “Coll. J. Faust/ Ankauf 1900”, “Typus”, nan), India and Myanmar. “Staatl. Museum für/ Tierkunde Dresden”, “Lectotype/ Material examined. CHINA. Yunnan: Xishuangbanna Parallelodemas perfectum/ Faust (Prena & Zhang/ de- Pref., Anma Xinzhai, 26.iv.2009 [#1941164], Meng (IZ- sign. 2014)” (SNSD). Paralectotypes 1 (female), Carin CAS 1); Xishuangbanna Pref., Menglun, Highway G213, Cheba, 900–1100 m, V XII.1888 (MSNG). Morimoto 21.xi.2009 [#1500895–96]; 22.xi.2009 [#1500894], Tang and Yoshihara (1996: 17), designation as type species Guo & Yao Zhiyuan (IZCAS 3); Xishuangbanna Pref., for Parallelodemas. Menglun, 12.iv.1994 [#1500897], Chen Xiaolin (IZCAS 1); Xishuangbanna Pref., Meng’a, 6.vi.1958 [#1799544– Diagnosis. Only two specimens of the sexually dimorphic 45], Wang Shuyong (IZCAS 2); Cangyuan County, Ban- P. perfectum are known, one of each gender. The species lao township, Dongnanhai, 2.v.2011 [#1941163], Huang forms a complex with P. feae, P. tardum, P. tumens and Xinlei (IZCAS 1). INDIA. Sikkim: Gopaldhara, Rung- P. vicinum, all of which have flush eyes, thoracic - ven bong Valley, H. Stevens (BMNH 3). West Bengal: Nam- trites with undivided setae and a male mesotarsus with su River, 21.vi.1918, H. Stevens (BMNH 1); Nurbong outward-directed clavate setae (Fig. 6). The profemur of [Estate], 1910s, W. K. Webb (BMNH 1). MYANMAR. the male P. perfectum is ventrally more expanded than Kachin: Teinthaw, V/1886, L. Fea (MSNG 1). Kayah: in the other species. Parallelodemas feae has ventrally

dez.pensoft.net 116 Jens Prena & Runzhi Zhang: A taxonomic revision of Parallelodemas Faust from South China...

and perpendicular to long axis of collum, ramus obsolete (as P. impar, Fig. 26). Distribution. The species is known from one site in Myanmar. Material examined. MYANMAR. Kayin: Karen Hills [ca. 25–35 km NE of Taungoo], V/1888, L. Fea (MSNG 1, SNSD 1).

Discussion

Species of Parallelodemas display an unusually diverse and complex suite of deviant morphological structures. Several occur in only a few species, such as the enlarged third tarsite or the deeply split setae on the ventrites. Oth- ers are male-specific, such as the setal fringes on rostrum, scape and mesotarsus, or the tubercles found on the first ventrite of an undescribed species near P. dimetans. The structural heterogeneity is increased further by the api- cally exposed epistome that often is worn off in females but rarely in males. Most of these traits can be found also in other tropical weevils, particularly in Dryophthorinae Figures 21–27. Female genitalia of Parallelodemas, sperma- and Baridinae (Davis 2009; Anderson et al. 2014; Prena theca: 21. P. dimetans; 22. P. petilum; 23. P. plumosum; 24. P. et al. 2014), although not as accumulated as in Parallelo- setifrons; 25. P. docile; 26. P. impar; 27. P. feae. Not to scale. demas. Very little is known about their functions and the few available observations may not always be transfer- able to other species. hirsute femora; P. tardum has a shorter male rostrum; P. It is long-known that numerous weevil species lack tumens is smaller (<5 mm); P. vicinum usually has denser incisors on the mandible (Lacordaire 1865; Ting 1936; vestiture on the mesepisternum and a less curved female Günther 1938). In some cases, the mandible moves almost rostrum. The few available female P. perfectum and P. vertically rather than transversely opposed as in most oth- tardum could not be distinguished with confidence. er (Horn 1873; McClenahan 1904; Marshall 1945; Redescription. Length 7.0–7.8 mm, width 1.8–1.9 Morimoto 1962; Pelsue and O’Brien 2011). However, dis- mm; integument black; ventral side with undivided setae, tally diverging mandibles with a convex inner face and in- basic vestiture inconspicuous on pronotum and elytron, cisor-like structures on the outer face are uncommon. They white squamiform setae at base of elytral interstriae 3 occur in several Dryophthorinae, such as Cyrtotrachelus and 4, postmedially on interstriae 3–5, on dorsal apex of Schönherr, Macrocheirus Schönherr, Otidognathus La- mesepimeron, metepisternum, prosternum, ventral face of cordaire, Protocerius Schönherr, Rhinostomus Rafinesque pro- and mesofemora, dorsal face of metafemur and oc- and, among the Baridinae, in Parallelodemas and some casionally on basolateral angles of pronotum; eyes flush Pascoe (Casey 1922; Vaurie 1970; Morimoto with head contour; frons and base of rostrum glabrous; and Yoshihara 1996). Many Rhynchitinae () male rostrum 1.06× as long as pronotum, ventrally with- and some Cholini, Erirhinini, Tychiini and Platypodinae out setae, prorostrum 0.36× rostral length and slightly (Curculionidae) have similar exodontous mandibles (Ting spatulate in dorsal view, epistome very slightly notched, 1936; Hamilton 1990; Thompson 1992) but with normal antennal scape with long, cupreous setae, club 1.6× as interior incisors and decussate apices. Three functions have long as wide; female rostrum 1.02× as long as pronotum, been attributed to exodontous mandibles. Daanje (1964) prorostrum 0.58× rostral length; prosternum slightly tu- was the first to point out that they occur in Rhynchitinae mescent (male) or gradually sloping (female) in front of that pupate in soil but not in Attelabinae that pupate in coxa, basal lobe notched; pro- and mesofemora ventrally leaf rolls. He concluded that the exterior tooth supports the with slender (male) or squamiform (female) setae, male weevil’s emergence to the surface. Depending on the spe- profemur somewhat angularly produced ventrally; tarsus cies group, the teeth are sheared off after emergence of the with tarsite 3 relatively small and excised to basal third, or are retained in one or both sexes (Daanje 1964; tarsite 5 longer than 3 and distinctly protruding beyond Dieckmann 1974; Riedel 2014). A second function of the anterior margin of 3, male mesotarsus with moderately exterior tooth is its usage during the preparation of the leaf long, clavate, outward directed setae; penis 3.1× as long roll in some Rhynchitinae (Daanje 1964, 1975). A third as wide, apex roundly narrowed with slightly produced possible function is related to oviposition. Kissinger (in tip (as P. feae, Fig. 19), internal sac with short, complex litt., quoted by Vaurie 1970) conjectured that the exterior sclerite (double hook); spermatheca with nodulus long tooth might be used for making oviposition holes, by rip-

dez.pensoft.net Dtsch. Entomol. Z. 61 (2) 2014, 105–119 117 ping through fibrous tissue of the monocotyledonous host Atelicus Waterhouse, Diabathrarius Schönherr, Macro­ plant. Eberhard (1983) observed that female Rhinostomus Champion, Syarbis Pascoe and Viticis Lea, and barbirostris (Fabricius) chew the oviposition hole and then are greatly reduced in several others. An enlarged third withdraw the rostrum with a series of sharp jerks. The ex- tarsite is particularly common in African Dryphthorinae odontous mandible may serve during the latter action for (e.g., Belorhynus Guérin-Méneville, Ichthyopisthen Au- enlarging or cleaning the hole. While the mandibles of the rivillius, Korotyaevius Alonso-Zarazaga and Lyal) and above-mentioned dryophthorines have a thick, excavated American grass-associated Baridinae (e.g., Macrobaris, outer face (Vaurie 1970), they are thin and blade-like in Nertinus Voss, Trachymeropsis Champion). In many cas- the baridines. It is conceivable that female Parallelode- es, the enlargement of the third tarsite is accompanied by mas employ the outer face for ripping and cutting through a reduction of the fifth. Many if not all of these species plant tissue as suggested by Kissinger, perhaps on culms live on swaying parts of their host and it is perceiv- hollow inside or filled with pith, but chewing is necessary able that the adhesive strength of the tarsus is increased to at least initiate the oviposition hole. Our own observa- by enlarging the surface of individual tarsites. tions showed that baridines without incisors are primarily Although the structural diversity of these predominant- pollen feeders and use their mandibles like pincers. Ovi- ly tropical weevils is appealing for morphological and position behavior has not been documented so far for any behavioral studies, systematic fieldwork is greatly ham- baridine with exodontous mandibles. The worn epistome pered by the still prevailing paucity of taxonomical and of many female Parallelodemas may or may not be related ecological information. Even the functions of rather ubiq- to the aberrant morphology and movement of the mandi- uitous structures, like the “prosternal horns” (Davis 2009; ble. Even though the epistome of Rhinostomus species is Davis and Engel 2010) present in Anthribidae, Nemony- similarly exposed and unprotected as in Parallelodemas, chidae and the curculionid subfamilies Baridinae, Cono­ we found no sign of wear in female R. barbirostris and R. derinae, Curculioninae and , have remained niger (Drury). However, similar wear is apparent in fe- largely unknown or were interpreted as being generally male Apostasimerus serrirostris Boheman, a palm-associ- indicative for ritual fighting, even though the latter usage ated Neotropical baridine with straight incisor area. has been observed in just one species (Eberhard and Garcia Gender-specific setae or setal patches occur relatively 2000; own observations) while similar structures have frequently on tibiae and ventrites of weevils (Eberhard other functions (Lacordaire 1863, p. 5; Lesne 1899, p. 1983; Lyal 1993; Schat et al. 2007) but are less common 143; Wood 1969, p. 43; Daanje 1975, p. 288; Thompson on the rostrum. Short fringes or fuzzy patches occur for in- 1992, p. 869). It is our hope that increased taxonomic and stance in Datonychus Wagner, Mogulones Reitter (Dieck- biogeographic knowledge will stimulate interest among mann 1972), Metamasius Horn (Vaurie 1968, 1970) and local researchers to conduct their own research on these Pterocolus Say (Hamilton 1998). We noticed long setal fascinating aspects of weevil diversity. fringes, like those present on the ventral rostrum of some male Parallelodemas species, in male Acythopeus bar- batus Pascoe, Conoproctus longipes (Casey), C. quadri- Acknowledgments pustulatus (Fabricius), Mycterus barbirostris Pascoe and M. imberbis Lea (probably a synonym of the former; all Our foremost thanks are extended to the Chinese Academy Baridinae). Observations on the usage of these setae are of Sciences for supporting the first author with a one-year available only for one species of Rhinostomus, the “bottle grant for international senior scientists (2012T1S0025). brush weevil”. Eberhard (1983) described how male R. Ren Li, who initiated this collaboration, and Wang Zhi­ barbirostris gently wipe the rostrum on the female’s pro- liang were tireless hosts and indispensable in making notum and elytron thereby apparently pacifying or immo- arrangements. We also acknowledge the help provided bilizing their chosen mate. Because this did not explain by other institute staff and Yang Jiani (formerly Beijing the presence of setae on the dorsal side of the rostrum, School of Forestry, now at Yale School of Forestry and En- the author speculated that other sensory functions may be vironmental Studies), who translated most label data and involved. However, his observations provided evidence determined geographic coordinates. Access to specimens that the setae have a behavioral, mating-related function. was arranged by Eva Sprecher-Übersax (Basel), Joachim If and how this applies to the exterior setae on the fifth Willers and Johannes Frisch (Berlin), Dirk Ahrens (Bonn), mesotarsite of some male Parallelodemas species re- Olaf Jäger (Dresden), Andrea Hastenpflug-Vesmanis and mains unknown. Similar setae occur in other weevils on Damir Kovac (Frankfurt a. M.), Maria Tavano and Ro- the interior (proximal) face of the basal three tarsites or berto Poggi (Genova), Karla Schneider, Joachim Händel all around, particularly on the protarsus, but not on the and Frank Steinheimer (Halle), Shep Myers (Honolulu), exterior face of the fifth. Max Barclay (London), Lutz Behne (Müncheberg), Bob The weevil tarsus typically is cryptopentamerous Anderson (Ottawa), Boris Korotyaev (St. Petersburg) (with a miniaturized fourth tarsite) but there are a few and Huang Junhao (Zhejiang). Bob Anderson and Ren Li exceptions and numerous modifications. The fourth and helped checking morphological characters of some spe- fifth tarsites are lost in species of genera such as Ano- cies. Open online access to this publication was sponsored plobaris Morimoto and Yoshihara, Anoplus Germar, by the Museum für Naturkunde, Berlin.

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References Lacordaire T (1865) Histoire naturelle des insectes. Genera des Coléoptères ou exposé méthodique et critique de tous les genres proposés jusqu’ici dans cet ordre d’insectes. Vol. 7. Roret, Paris Alonso-Zarazaga MA, Lyal CHC (1999) A world catalogue of families [1866], 620 pp. doi: 10.5962/bhl.title.67686 and genera of Curculionoidea (Insecta: Coleoptera) (Excepting Sco- Lesne P (1899) Sur l’usage des appendices mandibulaires caducs des lytidae and Platypodidae). Entomopraxis SCP, Barcelona. Brachyrrhinidae [Col.]. Bulletin de la Société Entomologique de Anderson RS, Marvaldi AE (2014) Dryophthorinae Schoenherr, 1825. France 68: 143–144. In: Leschen RAB, Beutel RG (Eds) Handbook of Zoology, Coleop- Lyal CHC (1993) Cryptorhynchinae (Insecta: Coleoptera: Curculionidae). tera, Vol. 3. DeGruyter, Berlin, 477–483. Fauna of New Zealand. Vol. 29. Manaaki Whenua Press, Lincoln. Casey TL (1922) Studies in the rhynchophorous subfamily Barinae of Lyal CHC, King T (1996) Elytro-tergal stridulation in weevils (Insecta: the Brazilian fauna. Memoirs on the Coleoptera 10: 1–520. Coleoptera: Curculionoidea). Journal of Natural History 30: 703– Daanje A (1964) Über die Ethologie und Blattrolltechnik von Deporaus 773. doi: 10.1080/00222939600770391 betulae L. und ein Vergleich mit den anderen blattrollenden Rhyn- Marshall GAK (1945) On Trephognathus: a new genus of African Bari- chitinen und Attelabinen-Coleoptera, Attelabinae. Verhandelingen dinae (Col. Curc.). Proceedings of the Royal Entomological Society der Koninklijke Nederlandse Akademie van Wetenschappen, Af- of London (B) 14: 21–24. doi: 10.1111/j.1365-3113.1945.tb00005.x deeling Natuurkunde 56: 1–215. McClenahan EM (1904) The development of the rostrum in the rhyncho- Daanje A (1975) Some special features of the leaf-rolling technique of phorous Coleoptera. Psyche 11: 89–102. doi: 10.1155/1904/95750 Byctiscus populi L. (Coleoptera: Rhynchitini). Behaviour 53: 285– Morimoto K (1961) On new Curculionidae from Japan (Coleoptera). 316. doi: 10.1163/156853975x00236 Kontyû 29: 22–27. Davis SR (2009) Morphology of Baridinae and related groups (Coleop- Morimoto K (1962) Comparative morphology and phylogeny of the su- tera, Curculionidae). ZooKeys 10: 1–136. doi: 10.3897/zookeys.10.47 perfamily Curculionidae of Japan (Comparative morphology, phylog- Davis SR, Engel MS (2010) Antiquity and evolution of prosternal horns eny and systematics of the superfamily Curculionidea of Japan. I). in baridine weevils (Coleoptera: Curculionidae). Journal of Paleon- Journal of the Faculty of Agriculture, Kyūshū University 11: 331–373. tology 84: 918–926. doi: 10.1666/09-160.1 Morimoto K, Yoshihara K (1996) On the genera of the Oriental Baridi- Dieckmann L (1972) Beiträge zur Insektenfauna der DDR: Coleoptera – nae (Coleoptera, Curculionidae). Esakia 36: 1–59. Curculionidae: Ceutorhynchinae. Beiträge zur Entomologie 22: 3–128. Oberprieler RG, Anderson RS, Marvaldi AE (2014) Curculionoidea La- Dieckmann L (1974) Beiträge zur Insektenfauna der DDR: Coleoptera – treille, 1802: Introduction, Phylogeny. In: Leschen RAB, Beutel RG Curculionidae (Rhinomacerinae, Rhynchitinae, Attelabinae, Apode- (Eds) Handbook of Zoology, Coleoptera, Vol. 3. DeGruyter, Berlin, rinae). Beiträge zur Entomologie 24: 5–54. 285–300. Eberhard WG (1983) Behavior of adult bottle brush weevils (Rhino­ Pelsue FW Jr, O’Brien CW (2011) A redefinition of the Curculionini of stomus barbirostris) (Coleoptera: Curculionidae). Revista Biologia the world, with a key to subtribes and genera, and two new genera: Tropicales 31: 233–244. Pseudoculio and Megaoculis (Coleoptera: Curculionidae: Curculi- Eberhard WG, Garcia-C JM (2000). Ritual jousting by horned Pariso- oninae). Zootaxa 3102: 27–49. schoenus expositus weevils (Coleoptera, Curculionidae, Baridinae). Prena J, Colonnelli E, Hespenheide HA (2014) Schoen- Psyche 103: 55–84. doi: 10.1155/2000/16361 herr, 1833. In: Leschen RAB, Beutel RG (Eds) Handbook of Zoolo- Faust J (1894) Viaggio di Leonardo Fea in Birmania e regioni vicine. gy, Coleoptera, Vol. 3. DeGruyter, Berlin, 577–589. LX. Curculionidae. Annali del Museo civico di storia naturale di Riedel A (2014) Attelabidae Billberg, 1820. In: Leschen RAB, Beutel Genova 34: 153–370. doi: 10.5962/bhl.title.48832 RG (eds). Handbook of Zoology, Coleoptera, Vol. 3. DeGruyter, Fea L (1896) Quattro anni fra i Birmani e le tribù limitrofe. Hoepli, Berlin, 328–355. Milano. Schat M, Sing SE, Peterson RKD (2007) External rostral characters for Günther K (1938) Revision der Gattung Otidognathus Lac. (Coleoptera differentiation of characters in the biological control agent Mecinus Curculionidae Calandrinae). Temminckia 3: 45–108. janthinus (Coleoptera: Curculionidae). Canadian Entomologist 139: Hamilton RW (1990) A revision of the weevil genus Eugnamptus 354–357. doi: 10.4039/n06-033 Schoenherr (Coleoptera: ) in America north of Mexico. Thompson RT (1992) Observations on the morphology and clas- Transactions of the American Entomological Society 115: 475–502. sification of weevils (Coleoptera, Curculionidae) with a key Hamilton RW (1998) Taxonomic revision of the New World Pteroco- to major groups. Journal of Natural History 26: 835–891. doi: linae (Coleoptera: Rhynchitidae). Transactions of the American En- 10.1080/00222939200770511 tomological Society 124: 203–269. Ting PC (1936) The mouth parts of the coleopterous group Rhyncho- Horn GH (1873) Contributions to a knowledge of the Curculionidae of phora. Microentomology 1: 93–114. the United States. Proceedings of the American Philosophical Society Vaurie P (1968) New weevils of the genus Metamasius from Central 13: 407–469. and South America (Coleoptera, Curculionidae, Rhynchophorinae). Korotyaev BA (2002) The weevil tribe Neosharpiini Hoffmann belongs Novitates Zoologicae 2316: 1–10. in the subfamily Baridinae (Coleoptera: Curculionidae). Zoosys- Vaurie P (1970) Weevils of the tribe Sipalini (Coleoptera, Curculionidae, tematica Rossica 11: 192. Rhynchophorinae) Part 1. The genera Rhinostomus and Yuccaborus. Lacordaire T (1863) Histoire naturelle des insectes. Genera des American Museum Novitates 2419: 1–57. Coléoptères ou exposé méthodique et critique de tous les genres Vinciguerra D (1890) Viaggio di Leonardo Fea in Birmania e regioni proposés jusqu’ici dans cet ordre d’insectes. Vol. 6. Roret, Paris, vicine. XIV. Pesci. Annali del Museo civico di storia naturale di 637 pp. doi: 10.5962/bhl.title.67686 Genova 9: 129–362, 1 map, plates VII–XI.

dez.pensoft.net Dtsch. Entomol. Z. 61 (2) 2014, 105–119 119

Voss E (1937) Ein weiterer Beitrag zur Kenntnis der Curculioniden Javas. burg. Nachtrag zum IX. Teil Insecta VI. Mitteilungen aus dem Ham- Tijdschrift voor Entomologie 80: 127–166. burgischen Zoologischen Museum und Institut 76: 395–468. Voss E (1941) Bemerkenswerte und unbeschriebene Rüsselkäfer aus Wood SL (1969) Additions to the horned bark beetle genus Cactopinus China und Japan (Col., Curc.). (92. Beitrag zur Kenntnis der Cur- Schwarz (Scolytidae). Coleopterists Bulletin 23: 42–51. culioniden). Mitteilungen der Münchener Entomologischen Ge- Yoshihara K, Morimoto K (1994) A new genus of the subfamily Bari- sellschaft 31: 887–902. dinae (Coleoptera, Curculionidae) from East Asia. Japanese Journal Voss E (1956) Über einige in Fukien (China) gesammelte Rüßler, V, of Entomology 62: 723–729. nebst einer neuen Gattung und Art aus Yunnan. (Col., Curc.) 115. Zherikhin VV, Gratshev VG (1995) A comparative study of the hind wing Beitrag zur Kenntnis der Curculioniden. Entomologische Blätter 51 venation of the superfamily Curculionoidea, with phylogenetic implica- [1955]: 21–45. tions. In: Pakaluk J, Ślipiński SA (Eds) Biology, Phylogeny, and Clas- Weidner H (1979) Die Entomologischen Sammlungen des Zoolo- sification of Coleoptera: Papers celebrating the 80th Birthday of Roy gischen Instituts und Zoologischen Museums der Universität Ham- A. Crowson. Muzeum i Instytut Zoologii PAN, Warszawa, 634–777.

dez.pensoft.net